Introduction

Indonesia is composed of 17,110 islands stretching 5,200 km in the southwestern Pacific and eastern Indian Oceans. The country embraces a total land area of over 5 million sq km, about 1 million sq km greater than the total land area of the United States. Indonesia spans two biogeographic zones, the Oriental and the Australian, and two floristic subregions, the Malesian and the Papuan (Takhtajan 1986). It encompasses at least 47 distinct natural ecosystems ranging from spectacular coral reefs, coastal mangrove swamps and lowland dry or humid forests, to deserts, montane rain forest, alpine meadows and glaciers. Although covering only about 1.3% of the earth's surface, Indonesia is one of the world's richest areas for biodiversity possessing approximately 17% of the total number of recorded species in the world. Contained within its boundaries are 17% of the world's bird species; 16% of the world's amphibians and reptiles; 12% of the world's mammals; 37% of the world's fish; and 11% of the world's flora. Glaringly absent from all published accounts of Indonesian biodiversity are estimates of fungal diversity. Along with other microorganisms, the fungi of Indonesia have received little attention from the scientific community. Historical literature, exsiccati and recent fungal collections from the region suggest, however, that the mycota of Indonesia is equally as diverse, although mostly undocumented.
Biodiversity inventory is a high priority of the Government of Indonesia. To help fill the gap in our knowledge of Indonesian fungi we have initiated a project funded by the National Science Foundation in cooperation with the Lembaga Ilmu Pengetahuan Indonesia [LIPI - Indonesia Institute of Science] and Pusat Penelitiam Dan Pengembangan Biologi [PPPB] that is designed to document the diversity of Agaricales from two of the major islands in Indonesia, viz., Java and Bali.

Taxonomic Scope of the Project:

The Agaricales (sensu Singer, 1986) include all holobasidiomycetes with fleshy fruitbodies forming lamellate or smooth hymenophores [= agarics] or poroid hymenophores [= boletes], plus all reduced forms, secotioid and hypogeous taxa putatively derived from them. The order Agaricales represents a paraphyletic group of organisms at least 100 million years old categorized in three suborders (Agaricineae, Russulineae, Boletineae) containing 17 families composed of 230 genera with over 7000 species worldwide (Singer, 1986). Agarics and boletes serve numerous intrinsic ecological roles: 1) Many are saprotrophic organisms that play an extremely important role in nutrient retention and recycling in plant communities, as primary or secondary decomposers of organic material; 2) Others are ectomycorrhizal symbionts that are essential components of healthy plant communities, mediating mineral and water uptake, and often providing protection from disease and predation by soil microorganisms; 3) The extensive mycelium of agarics and boletes is important in soil genesis, and aids in binding soil and litter, thereby reducing the amount of erosion on steep slopes and the concomitant potential loss of available nutrients; and 4) The vegetative mycelium and fleshy reproductive structures of these fungi serve as a primary food source for the larval and adult stages of numerous arthropod taxa, molluscs, and for some vertebrates. Agarics and boletes are an integral component of all terrestrial ecosystems.
We hypothesize that a minimum of 500 species of Agaricales occur in Java and Bali, although the total number of species is probably closer to 700. It is difficult to estimate how many of these may represent new species, although 50% is not an unreasonable figure. In our recent monograph of a single genus (Marasmius) from a similar underexplored tropical region in southeastern Asia (Papua New Guinea), 14 of 34 taxa (41%) were new to science (Desjardin & Horak 1997). In a monograph of the agaric genus Astrosporina from Indomalaysia and Australasia (Horak 1979b), 21 of 30 species reported (70%) were new to science, while in a treatment of Entoloma from the same region, 165 of 234 taxa (70%) were new (Horak 1980d). Similarily, in a recent treatment of 17 genera of white-spored agarics from Malaysia (Corner 1994), 132 of 168 species (78%) were new to science. Hawksworth (1993) presented a table itemizing the percentage of new species reported from Malaysia by Corner between 1950 and 1991 in nine different groups of macromycetes. Out of a total of 786 species reported, 517 (66%) were described as new. Given these data, we contend that an estimate of 50% new taxa from Java and Bali is reasonable and probably an underestimate. All new species encountered during this project will be published in refereed scientific journals, and a comprehensive mycota including all known Agaricales from Java and Bali will be published in printed and electronic format at the end of the project.

Geographic Scope of the Project:

Although we realize the need for and value of a comprehensive mycota of all of Indonesia, there are three major reasons why we have chosen to restrict our project to only the islands of Java and Bali. First, Indonesia is too large and too geographically and ecologically diverse to sample in its entirety in a 4 year time frame. One cannot expect to generate a database that reflects a reasonable representation of the mycological diversity present in the region in that short period of time. It is more realistic to select and sample several smaller land masses within the boundaries of Indonesia from which a preliminary database of fungal diversity can be generated. Java and Bali are two major Indonesian islands along the intensely volcanic Ring of Fire, both part of the Oriental biogeographic zone to the west of the Wallace Line, and part of the Malesian floristic subregion. Java is about 660 km long by 60-200 km in width with a total area of about 132,000 sq km, whereas Bali is about 140 km long by 80 km wide with a total area of about 5620 sq km. Both islands support limited but locally extensive areas of primary forest with high diversity of fleshy fungi, and are of appropriate size for adequate samping within four years. A database of fungi from the Oriental biogeographic zone (Malesian floristic subregion) generated during this project can then be compared with our data from the Australian biogeographic zone (Papuan floristic subregion) to address questions concerning biogeography, evolutionary patterns and species dispersal mechanisms (e.g., Desjardin & Horak 1997; Horak 1970, 1971a-b, 1973, 1976a-b, 1979b-i, 1980a, d-h, 1983, 1988a, 1995; Horak & Desjardin 1994).
Second, minimal mycological exploration has been conducted in Indonesia, and consequently limited baseline data are currently available concerning the diversity of Agaricales. As indicated above, we expect to encounter 500-700 species of Agaricales in Java and Bali. Because so little prior mycological research has been conducted in the area, and because so many of the associated plant communities and plant species are unique, we expect many of the fungal species encountered will be undescribed (possibly as many as 50%). Our experience in pioneering mycological biotic survey research in similar underexplored tropical regions of the world supports our contention. We contend that an area the size of Java and Bali with the expected diversity of 500-700 species is manageable in the context of the proposed project.
Third and most significantly, nearly all of the historical mycological research in Indonesia was conducted in Java and Bali and most Indonesian species of agarics and boletes were collected on these two islands. Java is the type locality for most Indonesian Agaricales. Unfortunately, most of these epithets are represented today only by their protologues, by water color illustrations, or by poorly preserved herbarium specimens. Good quality voucher specimens from which accurate species circumscriptions can be generated (and from which DNA can be isolated) are nearly nonexistant. Species previously described from the region must be recollected, meticulously preserved and accurately described in order to generate a meaningful survey of fungal diversity. These extant epithets must be linked to quality specimens to stabilize their nomenclature and species circumscriptions. For these reasons it is logical and most efficient to limit the scope of the initial project to the sampling and documentation of the Agaricales of only two Indonesian islands, specifically Java and Bali. Some researchers may suggest that it is more productive to initiate a biotic survey of Indonesian fungi on more pristine islands such as Sumatra or Sulawesi. We contend that until the nomenclature and taxonomy of the hundreds of extant epithets is stabilized through recollecting and analyzing topotype material, one runs the risk of creating superfluous epithets and promoting folkloric taxonomic concepts.

Known Levels of Diversity:

Léveillé (1844) and Moritzi (1845-1846) were the first to describe agaric species from Java. The earliest listings of Indonesian Agaricales that we are aware of were published in 1922 by Overeem & Overeem-de Haas in which 12 boletes and 273 agarics were reported, appended by Boedijn (1940) with another 28 species. Of these 313 taxa, 228 were reported from Java and Bali (the remainder from other Indonesian islands). Our current knowledge of the occurrence and distribution of Indonesian Agaricales is a direct reflection of the peregrinations of systematic mycologists, and the serendipitous nature of encountering ephemeral basidiomes. There have been no comprehensive biotic surveys or long-term sampling of basidiomycetous fungi conducted anywhere in Indonesia. Since Overeem's and Boedijn's reports, 46 additional papers have been published in which Agaricales of Java and Bali have been reported (Boedijn 1951a-b, 1960; Corner 1954, 1966, 1970, 1972, 1974, 1981, 1994, 1996; Corner & Bas 1962; Desjardin 1995; Guzman 1983; Hennings 1894, 1899, 1900, 1901, 1903; Holtermann 1898; Horak 1976, 1977a b, 1978, 1979a-c, 1980a-d, 1981a, 1983, 1988a, 1989; Léveillé 1844; Moritzi 1845-1846; Overeem 1926, 1927; Patouillard 1987; Singer 1945a-b, 1974, 1989; Singer & Smith 1958; Wakker 1844; Zollinger 1844, 1854). Most of these publications are monographs of selected genera from the greater geographic area of Indomalaysia and Australasia, in which a few specimens collected by the authors during their travels through Java and Bali were reported. A total of 353 species belonging to 92 genera have been reported to date from Java and Bali. An itemization of these is provided on our web page entitled "Fungi Previously Reported From Indonesia."

Status of Existing Collections:

The most extensive and historically significant collection of Indonesian Agaricales resides at Herbarium Bogoriense (BO), Bogor, Java. During a recent research trip to Bogor, we observed 1637 specimens deposited there, almost all of which were collected between 1919 and 1937 by the Dutch mycologists C. van Overeem, K. B. Boedijn and W. M. Docters van Leeuwen. Over 98% of these specimens are preserved in 70% alcohol in old, wax-sealed jars of various sizes. The specimens are organized in the herbarium according to jar size. Minimal label data are glued to the face of most jars or are deposited within the jars. Of these 1637 specimens, 480 are determined only to order, 862 are determined only to genus (representing 70 genera), while only 295 are determined to species, representing 122 species. The taxonomy and nomenclature dates to the 1920's and 1930's. An electronic database of these specimens with data in 16 fields resides at BO. Descriptive notes or watercolor illustrations depicting the morphological features of the basidiomes that comprise each specimen are available for less than 2% of the specimens. Unfortunately, for most Agaricales it is nearly impossible to accurately identify specimens to species rank without adequate notes on basidiome morphology and ecology. Consequently, the collection at BO, although of significant historical value, is of limited use for initiating an accurate and representative database of Indonesian Agaricales. Until fresh material is collected, described, and used as taxonomic reference material, most of the specimens at BO will remain undeterminable. It is obvious from looking at the 1600 jars, however, that there is a great diversity of Agaricales in Indonesia.
Other significant exsiccati holdings are a few type specimens of Berkeley (at K), von Höhnel (at FH) and Léveillé (at P). Although Hennings (1900) was the most significant author of Indonesian Agaricales, all of his pertinent specimens were lost from Berlin during World War II. Material collected by Corner and included in his books on Malesian agarics [primarily from Malaya, Borneo and the Soloman Islands (Corner 1972, 1994, 1996)] will soon be available for study from Edinburgh. Until his recent death, all of Corner's specimens were housed in his private herbarium and unavailable for study. Many of Corner's specimens are preserved in formalin and ethanol, rendering them of little value for future molecular studies. The best quality specimens, numbering several hundred, are those collected by Horak (at ZT).

Urgency for the Project:

Java and Bali are careening towards ecological disaster. Java, about the size of New York state and only 7% of Indonesia's total land area, has a population that has tripled this century reaching over 110 million people. The population of its capital, Jakarta, presently at 10 million, is expected to double by the year 2020. Java's population density is around 850 people per sq km, more than twice that of England (Europe's most populated country) or Japan. Over 60% of Indonesia's population lives on Java. Likewise, Bali is one of the more populated countries in the world. On a land mass smaller than the state of Connecticut live nearly 3 million people. None of these figures include the thousands of tourists that visit these popular vacation destinations each year. Under the enormous pressure of population, deforestation is progressing at a frightening pace. East Java alone consumes 14 million cubic meters of firewood per annum, equal to two thirds the peak lumber export of both Sumatra and Kalimantan (Dalton 1995). Throughout Java and Bali, vast areas below 2000 m are devoid of primary forests. The few remaining areas of primary forest that support a large variety of fleshy fungi are primarily in National and State Parks and Reserves or on inaccessible (i.e., not suitable for agriculture or habitation) mountain slopes. The fleshy fungi of these few primary forests must be documented soon.
Along with exceedingly dense human population comes pollution. Jakarta and the surrounding industrial area (known as Jabotabek; population 17 million) is one of the more polluted megalopoli in the world, rivaled only by Mexico City. Although there have been recent attempts at government regulated control of air pollution, the goals have not been fully realized. Fleshy fungi, especially ectomycorrhizal taxa, are very sensitive to the effects of air pollution (lowered soil pH, heavy metals, hydrocarbon toxicity) and there is abundant evidence of declining fungal diversity in European forests as a direct result of air pollution (Arnolds 1985, 1988, 1991a, b; Cherfas 1991; Jansen & Dobben 1987; Jansen & de Vries 1988; Meyer 1984; Termorshuizen & Schaeffers 1987). One can predict a similar pattern of diversity loss as a result of poor air quality in Java and Bali, particularly on ectomycorrhizal taxa of primary forests near population centers. Of special concern is the mycologically important Gede Pangrango National Park and adjacent Cibodas Botanical Gardens (Kebun Raya Cibodas) within 100 km of the ever-expanding Jabotabek population center. This site will be intensively surveyed as a part of this project (see Collecting Sites ).

Project Support:

This four year project to document the diversity of Agaricales from Java and Bali is co funded by the Biotic Surveys and Inventories Program and the International Programs (East Asia and Pacific) of the National Science Foundation (Grant. No. DEB-9705083). In Indonesia, the project is conducted in cooperation with Lembaga Ilmu Pengetahuan Indonesia (LIPI - Indonesian Institute of Sciences) and Pusat Penelitian Dan Pengembangan Biologi (PPPB - Research and Development Centre for Biology). Our Indonesian sponsor and collaborator is Dr. Mien Rifai, Asst. Minister of State for Research and Technology Development. He is also the Principal Mycologist at Herbarium Bogoriense, Bogor, and author of numerous papers on Indonesian fungi.